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A New Interpretation of Dragonfly Wing Venation Based Upon Early Upper A new interpretation of dragonfly wing venation based upon Early Upper Carboniferous fossils from Argentina (Insecta: Odonatoidea) and basic character states in pterygote wings EDGARF. RIEK 19 DufJL Street, Ainslie, Conberrcl, A .C.T.,2602, Austrcllia AND JARMILAKUKALOVA-PECK' Department of Geology, Carleton Universih, Ottc~wcl,Ont.. Cnnndo K IS 5B6 Received May 4. 1983 RIEK,E. F., and J. KUKALOVA-PECK.1984. A new interpretation of dragontly wing venation based upon Early Upper Carboniferous fossils from Argentina (Insecta: Odonatoidea) and basic character states in pterygote wings. Can. J. Zool. 62: 1150- 1166. The oldest known odonatoid wings are described from the Namurian of Argentina: Eugeropteron lunatum Riek n.g. et sp. and Geropteron arcuatum Riek n.g. and sp. (Meganisoptera: Meganeurina: Eugeropteridae n, fam.). The wings are gener- alized and support a reinterpretation of the venation of living Odonata as being fully homologous to that of other pterygotes and closely related to Ephemeropteroidea, but different from Neoptera. Therefore, Paleoptera is a valid phylogenetic unit, and Odonatoidea and Ephemeropteroidea are sister groups. RIEK.E. F.. et J. KUKALOVA-PECK.1984. A new interpretation of dragonfly wing venation based upon Early Upper Carboniferous fossils from Argentina (Insecta: Odonatoidea) and basic character states in pterygote wings. Can. J. Zool. 62: 1 150- 1 166. On trouvera ici la description des plus vieilles ailes odonatoi'des jamais rencontrees: il s'agit d'ailes d'Eugeropteron lunatum n. gen. et n. sp. et de Geropteron arcuatum n. gen. et n. sp. Riek (Meganisoptera: Meganeurina: Eugeropteridae n. fam.) provenant du Namurien d'Argentine. Les ailes suivent le plan de base et donnent lieu a une reinterpretation de la nervation des Odonates rkcentes selon laquelle les ailes des Odonates recentes seraient entierement homologues des ailes d'autres ptkrygotes, trks rapprochees des ailes d'Ephemeropteroidea, mais differentes des ailes de Neoptera. L'appellation Paleoptera coiffe donc un ensemble phylogenktique valide et il est juste de considerer comme des groupes-soeurs les Odonatoidea et les Ephemeropteroidea. [Traduit par le journal] Introduction Pterygote wing articulation was recently studied by Most entomologists follow Martynov (1924) and agree Kukalova-Peck ( 1983), who compared modern and Paleozoic that the pterygotes are naturally divided into Paleoptera and dragonflies with modern and Paleozoic Ephemeroptera, Paleo- Neoptera (Hennig 198 1 ). Nevertheless, some consider Paleo- zoic Paleodictyopteroidea (Paleodictyoptera, Megasecoptera, ptera not to be a natural phylogenetic group, but rather a grade Diaphanopterodea, and Permothemistida (=~rchodonata~)), (Lemche 1940; Boudreaux 1979; Rohdendorf and Rasnitsyn and primitive modern Neoptera. Multiple comparisons of artic- 1980; Rasnitsyn 1980; Kristensen 198 1 ; Matsuda 1970, 198 1 ). ular systems in different evolutionary stages showed that the Sometimes Odonata are assumed to be closer to Neoptera than odonatoid articulation evolved in a way similar to that of other to the Ephemeroptera (Kristensen 198 I), while other times, the Paleoptera, but unlike that of Neoptera. wings and wing articulation of Odonata are judged to be so The position of odonatoids within the Paleoptera, namely different that insect wings are believed to have originated whether they are more closely related to the paleodicty- twice: in the pterygote ancestor and in the odonatan ancestor opteroids or to the ephemeropteroids, is not clearly shown in (La Greca 1980; Matsuda 198 1 ). the articulation (Kukalova-Peck 1983). However, the closer These last two unorthodox evolutionary ideas find support in relationship to ephemeroids is evident in the veinal characters, the erroneous belief that the odonate wing venation and wing if the odonatoid veinal system presently used is corrected ac- articulation are unique and cannot be fully homologized with cording to the clues found in the fossil record. the other pterygote orders. The goal of this paper is to provide documentation that the We contend that wings and wing articulation originated at odonatoid venational system does not lack any veins as is the same time and brought winged insects into existence as a presently believed, and that it shares the ground plan with the group; therefore, they are considered as the basic apomorphic ephemeroids . characters of the pterygotes. It is believed that the split into Paleoptera, Neoptera, and the major ordinal evolutionary lin- Odonatan wing venation eages was initiated mainly by the radiation of the pterygote The interpretation of odonatan wing venation has a long proto-wings into diverse flight-adaptive structures (Martynov history of disagreement. Comstock and Needham ( 1898, 1899) 1924; Kukalova-Peck 1978, 1983; Hennig 198 1 ). If these con- erroneously assumed that veins are formed around tracheae and clusions are correct and wings are monophyletic, then odonatan wing venation and articulation must be fully homologous to 'we follow Sinichenkova (1980) in calling the order Permo- that of the other pterygotes. themistida Martynov, 1938, rather than Archodonata Martynov. 1932, because the group belongs in the paleodictyopteroids and has 'Author to whom reprint requests should be addressed. no relationship to Odonata. RIEK AND KUKALOVA-PECK 1151 that the nymphal tracheation recapitulates the ancestral vena- of with R, and the radial vein as the plesiomorphic double tion of adults. In fact, the preveinal blood channels in juveniles radius of primitive Paleodictyoptera (Figs. 6, 7) and fossil are invisible but present, and tracheae penetrate into them and living Ephemeroptera, i.e., composed of two radial stems secondarily and sometimes quite erratically. An erroneous RA and RP, the odonatan venation will lose its "strange" char- tracheae-based veinal system was adopted for odonates by acter and will become related to Ephemeroptera and Paleo- Needham ( 1903). Owing to vagaries of tracheation, the wings dictyoptera. Both "missing" veins MP- and CuA+ will then were interpreted as being completely unrelated to those of other be present in the venation, and the scheme will be at last fully pterygotes, i.e., RP (Rs)' was a convex vein crossed by M, and homologous with that in the other pterygote orders. M2, etc. Later on, numerous attempts were made by a number The original, generalized odonatan veinal scheme is indi- of authors to provide a homologous veinal scheme, namely cated in many Paleozoic Meganisoptera, but especially in- by Lameere (1922), Tillyard (1914, 1915, 1916, 1917, 19220, formative are two highly generalized (and probably also 19226, 1925a, 19256, 1926, 1928, 1935), Tillyard and the oldest known) dragonfly wings, which were recently Fraser ( 1938 - 1940), Martynov ( 1924), Carpenter ( 1 93 1 , found in the Early Upper Carboniferous deposits of Argentina. 1938, 1939, 1943n, 19436, 1944, 1947, 1960), Forbes (1943), Their venation represents the "missing link" between the Zalessky ( 1944), Fraser ( 1938, 1948, 1957), Needham ( 1903, typical odonatoid pattern and that of the ephemero-odonatoid 1951), Hamilton (1971, 1972), and Carle (1982). The inter- common ancestor. The Argentinian fossil dragonflies are pretations differ considerably, but none of them has been found described in the following text and their venation is compared to be correct in comparison with the venation of the oldest and with fossil and modern odonates, ephemeropteroids, and most generalized fossil dragonflies, which are described in this paleodictyopteroids. paper. One reason why there are so many seemingly feasible interpretations of odonate venation lies in the fact that in mod- Occurrence of the fossils ern dragonflies the primary veins alternate with the secondary Two very well preserved and almost complete insect wings intercalar veins and an occasional veinal supplement, and can- were found near Malanzan, Cuesta de la Herradura, La Rioja, not be readily distinguished from each other. However, sec- Argentina, in the carbonaceous group of the La Divisoria ondary veins are not fully tracheated and are sometimes miss- Member of the Malanzan Formation, by Drs. S. Archangelsky ing in the more primitive fossil dragonflies. Another reason is and R. Leguizamon. The insects were in association with a the basal parallel alignment of the fused veinal stems, which in plant assemblage including Vqjnowsba argentina Archangel- modern dragonflies is accompanied by disappearance of the sky & Leguizamon , Ginkgophvllum diuzii Archangelsky & original veinal crossings between these stems. If the crossings Arrondo, and Botrychiopsis u)eissiana Kurtz, as well as a were present, they would indicate the pattern under which the number of as yet undescribed plant species (S. Archangelsky, original fusions took place. The contribution of the fossil personal communication). dragonflies introduced here is that they (i) preserve ancestral Previous records of this fossiliferous horizon mentioned a character states of venation devoid of most secondary veins, Devonian or Early Carboniferous age, whereas Archangelsky and (ii) show the original pattern of veinal fusions before the and Leguizamon ( 197 I) thought that the strata might be dated crossings disappeared. Thus, generalized fossil dragonflies near the Westphalian -Stephanian boundary. Azcuy ( 1975) give clues to how the venation evolved. referred the horizon to the Namurian (Early Upper Carbon- Presently, the most widely followed odonatan veinal scheme iferous) after a palynological
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